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Saturday, August 08, 2009

On the Origins of Eukaryotes

Carl Zimmer has an article in Science with a provocative title: On the Origin of Eukaryotes. This is well-timed since it appears just when I've returned from a meeting on this very topic. [Go here if you can't see the article on the Science website.]

One of the things we learned at the meeting is that the Woose tree of life is almost certainly an over-simplification at best and wrong at worst. It is no longer possible to claim that eukaryotes have a simple vertical descent relationship with any archaebacterium (or any bacterium, for that matter).

Instead, the early history of life is characterized by a web or a net involving multiple gene exchanges between all primitive species. After some time, the major divisions of life emerged from this "soup" and became separate lineages with an semi-independent history. This view dates back ten years or so and it's illustrated by a figure that Ford Doolittle published in the February 2000 issue of Scientific American. I've used this figure several times. Here it is again so you can see how it relates to Carl's article.

In the case of eukaryotes, the history is complicated by an endosymbiotic event where a proteobacterium was engulfed and evolved into mitochondria. That explains many of the eukaryotic genes with a clear bacterial origin. Those genes, can be reliably traced to a particular lineage of proteobacteria. What this shows is that by the time of the endosymbiosis most of the main lineages of prokaryotes had emerged from the soup and become fairly well-defined.

This doesn't explain the origins of the host cell. That cell presumably had some of the features of modern eukaryotes. Where did it come from? Was it part of an ancient lineage that formed during the gene exchange period of evolution suggesting that some eukaryotic features are ancient? Was it formed by a fusion between a primitive bacterial cell and a primitive archaebacterium? (Or, did archaebacterial arise from a fusion of a primitive eukaryotic cell and a primitive bacterium?)

Some people even believe that the ancient host progenitor of eukaryotes arose fairly late in the game and was only related to archaebacteria, either through a recent common ancestor or from an archaebaterial species within the archaebacterial clade. This is not consistent with the tree shown above but that's OK.

These two hypotheses on the archaebacterial origin of the host cell are the ones that Carl Zimmer highlights in his article: the Three Domain Tree and the Eocyte Tree.

I don't think either of these trees comes close to representing the true history of eukaryotic cells. I don't think it's even possible to represent that history by a tree. Zimmer mentions this possibility in passing but I don't think he does justice to the controversy over the tree of life.

The controversy is not just about which branch of the archaebacterial tree the eukaryotes came from. It's about whether they came from the archaebacerial lineage at all or whether it's even appropiate to be talking about lineages and trees at this stage of the history of life.

Shifting gears slightly, I'd like to bring up another subject. Here's what Carl writes near the end of his article.

Whatever the exact series of events turns out to be, eukaryotes triggered a biological revolution. Prokaryotes can generate energy only by pumping charged atoms across their membranes. That constraint helps limit their size. As prokaryotes grow in size, their volume increases much faster than their surface area. They end up with too little energy to power their cells. Eukaryotes, on the other hand, can pack hundreds of energy-generating mitochondria into a single cell. And so they could get big, evolving into an entirely new ecological niche.

This is a widely believed explanation for the adaptive value of mitochondria and internal membranes. But many species of bacteria have internal membranes and, in the case of photosynthetic species, those internal membranes are packed full of energy-producing proteins.

Why couldn't bacteria have evolved internal membranes in order to get around the size limitation if it was that big of a deal? I don't see anything special about mitochondrial that couldn't have just as easily been handled by infolding of the inner membrane.

3 comments
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I'm disappointed that Carl Zimmer seems to have completely ignored (or been unaware of) the wonderfully intricate (if not too intricate) Neomuran Hypothesis by Cavalier-Smith. [Very] long story short (there's a 2002 review that goes on for 56 pages in TC-S writing, ie equivalent to about ~100pages in normal English =P), an ancestor of Archaebacteria and Eukarya lost its outer wall; Archaea responded by creating a glycoprotein wall and adapting to extreme environments; Eukarya developed a sophisticated cytoskeleton which then enabled phagocytosis, which in turn allowed the early Eukaryotes to become large predators. There's a good (and relatively short) review in TC-S 2009 Int J Biochem & Cell Biol, so don't rely on my butchered summary there...

Of course, then there's also bacteria that are capable of some sort of phagocytosis like thing, and there's even prokaryotic endosymbionts of other prokaryotes! But -generally- speaking, cytosk, phagocytosis and endomembrane sys are primarily eukaryotic features. Although we've only sampled a miniscule insignificant percentage of prokaryotic biodiversity, so who knows what's really out there...

I don't like the soup view very much, as it implies a sort of shapeless primitiveness of non-large/popular-eukaryotic life. Microbial life is full of fascinating unique features and sophisticated solutions to life's problems. There's algae (Warnowiid dinos) with camera eyes entirely out of subcellular components: a retinoid and hyaline lens. There's ridiculous RNA editing systems like that of Tryp kDNA. There's creatures with the whackiest lifestyles imaginable. There's massive bacteria up to 800um with strange membrane invaginations+vesicles at the surface (that apparently haven't been investigated yet). Actually, if you zoom in a bit, perhaps it's safe to say that -every- creature has some strange fascinating story to tell. They are by far not a primordial prelude to 'Higher Life'.

Thus, even if you'd like to emphasise the potential impacts of LGT, can we perhaps use a tree that doesn't completely and utterly fuck up the eukaryotic lineages? It's fairly old, even predates the final collapse of Archaezoa. Furthermore, it exhibits the savage fallacy of the 'Crown Eukaryote' reasoning; ie that animals, land plants and fungi are some sort of advanced organisms presiding over 'lower' 'unicellular' eukaryotic life (multicellularity arose up to about 16 times independently among eukaryotes; not just once before fungi+plants+animals as the tree implies). Sorry, it's just a massive pet peeve of mine, can't let it slip uncommented...

But yeah, I agree that the mitochondria --> big argument is rather strange. Never heard of it before... besides, you had to get big in the first place to be able to phagocytose bacteria; thus the thinking there is rather backwards.

/rant

Signed,"Defender of Marginalised Eukaryotes" =P (not that they need any help, considering they have pretty much taken over the world already...)

Laurence A. Moran

Larry Moran is a Professor Emeritus in the Department of Biochemistry at the University of Toronto. You can contact him by looking up his email address on the University of Toronto website.

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The old argument of design in nature, as given by Paley, which formerly seemed to me to be so conclusive, fails, now that the law of natural selection has been discovered. We can no longer argue that, for instance, the beautiful hinge of a bivalve shell must have been made by an intelligent being, like the hinge of a door by man. There seems to be no more design in the variability of organic beings and in the action of natural selection, than in the course which the wind blows.Charles Darwin (c1880)Although I am fully convinced of the truth of the views given in this volume, I by no means expect to convince experienced naturalists whose minds are stocked with a multitude of facts all viewed, during a long course of years, from a point of view directly opposite to mine. It is so easy to hide our ignorance under such expressions as "plan of creation," "unity of design," etc., and to think that we give an explanation when we only restate a fact. Any one whose disposition leads him to attach more weight to unexplained difficulties than to the explanation of a certain number of facts will certainly reject the theory.

Charles Darwin (1859)Science reveals where religion conceals. Where religion purports to explain, it actually resorts to tautology. To assert that "God did it" is no more than an admission of ignorance dressed deceitfully as an explanation...

Quotations

The world is not inhabited exclusively by fools, and when a subject arouses intense interest, as this one has, something other than semantics is usually at stake.
Stephen Jay Gould (1982)
I have championed contingency, and will continue to do so, because its large realm and legitimate claims have been so poorly attended by evolutionary scientists who cannot discern the beat of this different drummer while their brains and ears remain tuned to only the sounds of general theory.
Stephen Jay Gould (2002) p.1339
The essence of Darwinism lies in its claim that natural selection creates the fit. Variation is ubiquitous and random in direction. It supplies raw material only. Natural selection directs the course of evolutionary change.
Stephen Jay Gould (1977)
Rudyard Kipling asked how the leopard got its spots, the rhino its wrinkled skin. He called his answers "just-so stories." When evolutionists try to explain form and behavior, they also tell just-so stories—and the agent is natural selection. Virtuosity in invention replaces testability as the criterion for acceptance.
Stephen Jay Gould (1980)
Since 'change of gene frequencies in populations' is the 'official' definition of evolution, randomness has transgressed Darwin's border and asserted itself as an agent of evolutionary change.
Stephen Jay Gould (1983) p.335
The first commandment for all versions of NOMA might be summarized by stating: "Thou shalt not mix the magisteria by claiming that God directly ordains important events in the history of nature by special interference knowable only through revelation and not accessible to science." In common parlance, we refer to such special interference as "miracle"—operationally defined as a unique and temporary suspension of natural law to reorder the facts of nature by divine fiat.
Stephen Jay Gould (1999) p.84

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My own view is that conclusions about the evolution of human behavior should be based on research at least as rigorous as that used in studying nonhuman animals. And if you read the animal behavior journals, you'll see that this requirement sets the bar pretty high, so that many assertions about evolutionary psychology sink without a trace.

Jerry Coyne
Why Evolution Is TrueI once made the remark that two things disappeared in 1990: one was communism, the other was biochemistry and that only one of them should be allowed to come back.

Sydney Brenner
TIBS Dec. 2000
It is naïve to think that if a species' environment changes the species must adapt or else become extinct.... Just as a changed environment need not set in motion selection for new adaptations, new adaptations may evolve in an unchanging environment if new mutations arise that are superior to any pre-existing variations

Douglas Futuyma
One of the most frightening things in the Western world, and in this country in particular, is the number of people who believe in things that are scientifically false. If someone tells me that the earth is less than 10,000 years old, in my opinion he should see a psychiatrist.

Francis Crick
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Sydney Brenner
An atheist before Darwin could have said, following Hume: 'I have no explanation for complex biological design. All I know is that God isn't a good explanation, so we must wait and hope that somebody comes up with a better one.' I can't help feeling that such a position, though logically sound, would have left one feeling pretty unsatisfied, and that although atheism might have been logically tenable before Darwin, Darwin made it possible to be an intellectually fulfilled atheist

Richard Dawkins
Another curious aspect of the theory of evolution is that everybody thinks he understand it. I mean philosophers, social scientists, and so on. While in fact very few people understand it, actually as it stands, even as it stood when Darwin expressed it, and even less as we now may be able to understand it in biology.

Jacques Monod
The false view of evolution as a process of global optimizing has been applied literally by engineers who, taken in by a mistaken metaphor, have attempted to find globally optimal solutions to design problems by writing programs that model evolution by natural selection.